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''Lycoperdon perlatum'', popularly known as the common
puffball Puffballs are a type of fungus featuring a ball-shaped fruit body that bursts on impact, releasing a cloud of dust-like spores when mature. Puffballs belong to the division Basidiomycota and encompass several genera, including '' Calvatia'', '' ...
, warted puffball, gem-studded puffball, wolf farts or the devil's snuff-box, is a species of
puffball Puffballs are a type of fungus featuring a ball-shaped fruit body that bursts on impact, releasing a cloud of dust-like spores when mature. Puffballs belong to the division Basidiomycota and encompass several genera, including '' Calvatia'', '' ...
fungus in the family
Agaricaceae The Agaricaceae are a family of basidiomycete fungi and include the genus ''Agaricus'', as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. Taxonomy The family Agaricaceae was publish ...
. A widespread species with a cosmopolitan distribution, it is a medium-sized puffball with a round fruit body tapering to a wide stalk, and dimensions of wide by tall. It is off-white with a top covered in short spiny bumps or "jewels", which are easily rubbed off to leave a netlike pattern on the surface. When mature it becomes brown, and a hole in the top opens to release
spores In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
in a burst when the body is compressed by touch or falling raindrops. The puffball grows in fields, gardens, and along roadsides, as well as in grassy clearings in woods. It is edible when young and the internal flesh is completely white, although care must be taken to avoid confusion with immature fruit bodies of
poisonous Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...
''
Amanita The genus ''Amanita'' contains about 600 species of agarics, including some of the most toxic known mushrooms found worldwide, as well as some well-regarded edible species. This genus is responsible for approximately 95% of the fatalities result ...
'' species. ''L. perlatum'' can usually be distinguished from other similar puffballs by differences in surface texture. Several chemical compounds have been isolated and identified from the fruit bodies of ''L. perlatum'', including
sterol Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the g ...
derivatives, volatile compounds that give the puffball its flavor and odor, and the unusual
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
lycoperdic acid.
Extract An extract is a substance made by extracting a part of a raw material, often by using a solvent such as ethanol, oil or water. Extracts may be sold as tinctures, absolutes or in powder form. The aromatic principles of many spices, nuts ...
s of the puffball have
antimicrobial An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ...
and
antifungal An antifungal medication, also known as an antimycotic medication, is a pharmaceutical fungicide or fungistatic used to treat and prevent mycosis such as athlete's foot, ringworm, candidiasis (thrush), serious systemic infections such as cry ...
activities.


Taxonomy

The species was first described in the scientific literature in 1796 by mycologist
Christiaan Hendrik Persoon Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Linnaeus' mushroom taxonomy. Early life Persoon was born in South Africa at the Cape of Good Hope, the third child of an immi ...
.
Synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are ...
include ''Lycoperdon gemmatum'' (as described by August Batsch in 1783); the
variety Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
''Lycoperdon gemmatum'' var. ''perlatum'' (published by Elias Magnus Fries in 1829); ''Lycoperdon bonordenii'' (
George Edward Massee George Edward Massee (20 December 1845 – 16 February 1917) was an English mycologist, plant pathologist, and botanist. Background and education George Massee was born in Scampston, East Riding of Yorkshire, the son of a farmer. He was educat ...
, 1887); and ''Lycoperdon perlatum'' var. ''bonordenii'' (A.C. Perdeck, 1950). ''L. perlatum'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
of the genus ''Lycoperdon''.
Molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioch ...
analyses suggest a close
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
relationship with '' L. marginatum''. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''perlatum'' is
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
for "widespread". It is commonly known as the common puffball, the gem-studded puffball (or gemmed puffball), the warted puffball, or the devil's snuff-box;
Samuel Frederick Gray Samuel Frederick Gray (10 December 1766 – 12 April 1828) was a British botanist, mycologist, and pharmacologist. He was the father of the zoologists John Edward Gray and George Robert Gray. Background He was the son of Samuel Gray, a Londo ...
called it the pearly puff-ball in his 1821 work ''A Natural Arrangement of British Plants''. Because some
indigenous peoples Indigenous peoples are culturally distinct ethnic groups whose members are directly descended from the earliest known inhabitants of a particular geographic region and, to some extent, maintain the language and culture of those original people ...
believed that the spores caused blindness, the puffball has some local names such as "blindman's bellows" and "no-eyes".


Description

The fruit body ranges in shape from pear-like with a flattened top, to nearly spherical, and reaches dimensions of wide by tall. It has a stem-like base, and is whitish before browning in age. The outer surface of the fruit body (the exoperidium) is covered in short cone-shaped spines that are interspersed with granular warts. The spines, which are whitish, gray, or brown, can be easily rubbed off, and leave
reticulate Reticulation is a net-like pattern, arrangement, or structure. Reticulation or Reticulated may refer to: * Reticulation (single-access key), a structure of an identification tree, where there are several possible routes to a correct identificat ...
pock marks or scars after they are removed. The base of the puffball is thick, and has internal chambers. It is initially white, but turns yellow, olive, or brownish in age. The reticulate pattern resulting from the rubbed-off spines is less evident on the base. In maturity, the exoperidium at the top of the puffball sloughs away, revealing a pre-formed hole (
ostiole An ''ostiole'' is a small hole or opening through which algae or fungi release their mature spores. The word is a diminutive of "ostium", "opening". The term is also used in higher plants, for example to denote the opening of the involuted ...
) in the endoperidium, through which the spores can escape. In young puffballs, the internal contents, the
gleba Gleba (, from Latin ''glaeba, glēba'', "lump") is the fleshy spore-bearing inner mass of certain fungi such as the puffball or stinkhorn. The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The conti ...
, is white and firm, but turns brown and powdery as the
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s mature. The gleba contains minute chambers that are lined with
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others som ...
(the fertile, spore-bearing tissue); the chambers collapse when the spores mature. Mature puffballs release their powdery spores through the ostiole when they are compressed by touch or falling raindrops. A study of the spore release mechanism in ''L. pyriforme'' using high-speed
schlieren photography Schlieren photography is a process for photographing fluid flow. Invented by the German physicist August Toepler in 1864 to study supersonic motion, it is widely used in aeronautical engineering to photograph the flow of air around objects. ...
determined that raindrops of 1 mm diameter or greater, including rain drips from nearby trees, were sufficient to cause spore discharge. The puffed spores are ejected from the ostiole at a velocity of about 100 cm/second to form a centimeter-tall cloud one-hundredth of a second after impact. A single puff like this can release over a million spores. The spores are spherical, thick-walled, covered with minute spines, and measure 3.5–4.5  μm in diameter. The capillitia (threadlike filaments in the gleba in which spores are embedded) are yellow-brown to brownish in color, lack septae, and measure 3–7.5 μm in diameter. The
basidia A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly- ...
(spore-bearing cells) are club-shaped, four-spored, and measure 7–9 by 4–5 μm. The basidia bear four slender
sterigmata In biology, a sterigma (pl. sterigmata) is a small supporting structure. It commonly refers to an extension of the basidium (the spore-bearing cells) consisting of a basal filamentous part and a slender projection which carries a spore at the ti ...
of unequal length ranging from 5–10 μm long. The surface spines are made of chains of pseudoparenchymatous
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
e (resembling the
parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word π ...
of higher plants), in which the individual hyphal cells are spherical to elliptical in shape, thick-walled (up to 1 μm), and measure 13–40 by 9–35 μm. These hyphae do not have
clamp connection A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of Basidiomycetes fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rece ...
s.


Edibility

''Lycoperdon perlatum'' is considered to be a good
edible mushroom Edible mushrooms are the fleshy and edible fruit bodies of several species of macrofungi (fungi which bear fruiting structures that are large enough to be seen with the naked eye). They can appear either below ground (hypogeous) or above ground ...
when young, when the gleba is still homogeneous and white. They have been referred to as "poor man's sweetbread" due to their texture and flavor. The fruit bodies can be eaten after slicing and frying in batter or egg and breadcrumbs, or used in soups as a substitute for
dumpling Dumpling is a broad class of dishes that consist of pieces of dough (made from a variety of starch sources), oftentimes wrapped around a filling. The dough can be based on bread, flour, buckwheat or potatoes, and may be filled with meat, ...
s. As early as 1861, Elias Fries recommended them dried and served with salt, pepper, and oil. The puffballs become inedible as they mature: the gleba becomes yellow-tinged then finally develops into a mass of powdery olive-green spores. ''L. perlatum'' is one of several edible species sold in markets in the Mexican states of
Puebla Puebla ( en, colony, settlement), officially Free and Sovereign State of Puebla ( es, Estado Libre y Soberano de Puebla), is one of the 32 states which comprise the Federal Entities of Mexico. It is divided into 217 municipalities and its cap ...
and
Tlaxcala Tlaxcala (; , ; from nah, Tlaxcallān ), officially the Free and Sovereign State of Tlaxcala ( es, Estado Libre y Soberano de Tlaxcala), is one of the 32 states which comprise the Federal Entities of Mexico. It is divided into 60 municipaliti ...
. The fruit bodies are appealing to other animals as well: the
northern flying squirrel The northern flying squirrel (''Glaucomys sabrinus'') is one of three species of the genus '' Glaucomys'', the only flying squirrels found in North America.Walker EP, Paradiso JL. 1975. ''Mammals of the World''. Baltimore: Johns Hopkins Universi ...
(''Glaucomys sabrinus'') includes the puffball in their diet of non-
truffle A truffle is the Sporocarp (veggie), fruiting body of a subterranean ascomycete fungus, predominantly one of the many species of the genus ''Tuber (fungus), Tuber''. In addition to ''Tuber'', many other genera of fungi are classified as truf ...
fungi, while the "puffball beetle" ''
Caenocara subglobosum ''Caenocara'' is a genus of beetles in the family Ptinidae. Members of this genus are sometimes called puffball beetles. Species Species include: Nutritional analysis indicates that the puffballs are a good source of
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respon ...
,
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ...
s,
fat In nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such compounds, most commonly those that occur in living beings or in food. The term often refers specifically to triglycerides (triple es ...
s, and several
micronutrient Micronutrients are essential dietary elements required by organisms in varying quantities throughout life to orchestrate a range of physiological functions to maintain health. Micronutrient requirements differ between organisms; for example, huma ...
s. The predominant
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, ...
s in the puffball are
linoleic acid Linoleic acid (LA) is an organic compound with the formula COOH(CH2)7CH=CHCH2CH=CH(CH2)4CH3. Both alkene groups are ''cis''. It is a fatty acid sometimes denoted 18:2 (n-6) or 18:2 ''cis''-9,12. A linoleate is a salt or ester of this acid. L ...
(37% of the total fatty acids),
oleic acid Oleic acid is a fatty acid that occurs naturally in various animal and vegetable fats and oils. It is an odorless, colorless oil, although commercial samples may be yellowish. In chemical terms, oleic acid is classified as a monounsaturated om ...
(24%),
palmitic acid Palmitic acid (hexadecanoic acid in IUPAC nomenclature) is a fatty acid with a 16-carbon chain. It is the most common saturated fatty acid found in animals, plants and microorganisms.Gunstone, F. D., John L. Harwood, and Albert J. Dijkstra. The ...
(14.5%), and
stearic acid Stearic acid ( , ) is a saturated fatty acid with an 18-carbon chain. The IUPAC name is octadecanoic acid. It is a waxy solid and its chemical formula is C17H35CO2H. Its name comes from the Greek word στέαρ "''stéar''", which means tall ...
(6.4%). The immature 'buttons' or 'eggs' of deadly ''
Amanita The genus ''Amanita'' contains about 600 species of agarics, including some of the most toxic known mushrooms found worldwide, as well as some well-regarded edible species. This genus is responsible for approximately 95% of the fatalities result ...
'' species can be confused with puffballs. This can be avoided by slicing fruit bodies vertically and inspecting them for the internal developing structures of a mushroom, which would indicate the poisonous ''Amanita''. Additionally, amanitas will generally not have "jewels" or a bumpy external surface. The spores' surfaces have many microscopic spines and can cause severe irritation of the lung ( lycoperdonosis) when inhaled. This condition has been reported to afflict dogs that play or run where the puffballs are present.


Similar species

There are several other puffball species with which ''L. perlatum'' might be confused. '' L. nettyanum'', found in the
Pacific Northwest The Pacific Northwest (sometimes Cascadia, or simply abbreviated as PNW) is a geographic region in western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though ...
region of the United States, is covered in granular patches, but these granules adhere more strongly to the surface than those of ''L. perlatum''. '' L. pyriforme'' lacks prominent spines on the surface, and grows on rotting wood—although if growing on buried wood, it may appear to be terrestrial. The widely distributed and common '' L. umbrinum'' has spines that do not leave scars when rubbed off, a gleba that varies in color from dark brown to purple-brown at maturity, and a purple-tinged base. The small and rare species '' L. muscorum'' grows in deep moss. '' L. peckii'' can be distinguished from ''L. pyriforme'' by the lavender-tinged spines it has when young. '' L. rimulatum'' has purplish spores, and an almost completely smooth exoperidium. '' L. excipuliforme'' is larger and grayer, and, in mature individuals, the upper portion of its fruit body breaks down completely to release its spores. In the field, '' L. marginatum'' is distinguished from ''L. perlatum'' by the way in which the spines are shed from the exoperidium in irregular sheets.


Ecology and distribution

A
saprobic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ( ...
species, ''Lycoperdon perlatum'' grows solitarily, scattered, or in groups or clusters on the ground. It can also grow in
fairy ring A fairy ring, also known as fairy circle, elf circle, elf ring or pixie ring, is a naturally occurring ring or arc of mushrooms. They are found mainly in forested areas, but also appear in grasslands or rangelands. Fairy rings are detectable by ...
s. Typical habitats include woods, grassy areas, and along roads. It has been reported from ''
Pinus patula ''Pinus patula'', commonly known as patula pine, spreading-leaved pine, or Mexican weeping pine, and in Spanish as ''pino patula'' or ''pino llorón'', (''patula'' Latin = “spreading”) is a tree native to the highlands of Mexico. It grows f ...
'' plantations in
Tamil Nadu Tamil Nadu (; , TN) is a state in southern India. It is the tenth largest Indian state by area and the sixth largest by population. Its capital and largest city is Chennai. Tamil Nadu is the home of the Tamil people, whose Tamil languag ...
, India. The puffball sometimes confuses golfers because of its resemblance to a golf ball when viewed from a distance. A widespread species with an almost cosmopolitan distribution, it has been reported from Africa (
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
, Rwanda,
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
), Asia (China, Himalayas, Japan, southern India Iran), Australia, Europe, New Zealand, and South America (Brazil). It has been collected from subarctic areas of
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland is ...
, and
subalpine Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial f ...
regions in
Iceland Iceland ( is, Ísland; ) is a Nordic island country in the North Atlantic Ocean and in the Arctic Ocean. Iceland is the most sparsely populated country in Europe. Iceland's capital and largest city is Reykjavík, which (along with its ...
. In North America, where it is considered the most common puffball species, it ranges from
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S ...
to Mexico, although it is less common in Central America. The species is popular on postage stamps, and has been depicted on stamps from Guinea, Paraguay, Romania, Sierra Leone, and Sweden. The puffball bioaccumulates heavy metals present in the soil, and can be used as a
bioindicator A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
of soil pollution by heavy metals and
selenium Selenium is a chemical element with the symbol Se and atomic number 34. It is a nonmetal (more rarely considered a metalloid) with properties that are intermediate between the elements above and below in the periodic table, sulfur and telluriu ...
. In one 1977 study, samples collected from grassy areas near the side of an
interstate highway The Dwight D. Eisenhower National System of Interstate and Defense Highways, commonly known as the Interstate Highway System, is a network of controlled-access highways that forms part of the National Highway System in the United States. T ...
in
Connecticut Connecticut () is the southernmost state in the New England region of the Northeastern United States. It is bordered by Rhode Island to the east, Massachusetts to the north, New York to the west, and Long Island Sound to the south. Its cap ...
were shown to have high concentrations of
cadmium Cadmium is a chemical element with the symbol Cd and atomic number 48. This soft, silvery-white metal is chemically similar to the two other stable metals in group 12, zinc and mercury. Like zinc, it demonstrates oxidation state +2 in most of ...
and
lead Lead is a chemical element with the Symbol (chemistry), symbol Pb (from the Latin ) and atomic number 82. It is a heavy metals, heavy metal that is density, denser than most common materials. Lead is Mohs scale of mineral hardness#Intermediate ...
. ''L. perlatum''
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms biom ...
has been shown experimentally to remove mercury ions from
aqueous solution An aqueous solution is a solution in which the solvent is water. It is mostly shown in chemical equations by appending (aq) to the relevant chemical formula. For example, a solution of table salt, or sodium chloride (NaCl), in water would ...
s, and is being investigated for potential use as a low-cost,
renewable A renewable resource, also known as a flow resource, is a natural resource which will replenish to replace the portion depleted by usage and consumption, either through natural reproduction or other recurring processes in a finite amount of ti ...
, biosorptive material in the treatment of water and
wastewater Wastewater is water generated after the use of freshwater, raw water, drinking water or saline water in a variety of deliberate applications or processes. Another definition of wastewater is "Used water from any combination of domestic, industri ...
containing mercury.


Chemistry

Lycoperdic acid is an amino acid known only from ''L. perlatum''. Several
steroid A steroid is a biologically active organic compound with four rings arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes that alter membrane fluidity; and ...
derivatives have been isolated and identified from fruit bodies of ''L. perlatum'', including (''S'')-23-hydroxylanostrol,
ergosterol Ergosterol (ergosta-5,7,22-trien-3β-ol) is a sterol found in cell membranes of fungi and protozoa, serving many of the same functions that cholesterol serves in animal cells. Because many fungi and protozoa cannot survive without ergosterol, the ...
α-endoperoxide, ergosterol 9,11-dehydroendoperoxide and (23''E'')-lanosta-8,23-dien-3β,25-diol. The compounds 3-octanone, 1-octen-3-ol, and (''Z'')-3-octen-1-ol are the predominant components of the volatile chemicals that give the puffball its odor and flavor. Extracts of the puffball contain relatively high levels of
antimicrobial An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ...
activity against laboratory cultures of the human
pathogenic bacteria Pathogenic bacteria are bacteria that can cause disease. This article focuses on the bacteria that are pathogenic to humans. Most species of bacteria are harmless and are often beneficial but others can cause infectious diseases. The number of th ...
''
Bacillus subtilis ''Bacillus subtilis'', known also as the hay bacillus or grass bacillus, is a Gram-positive, catalase-positive bacterium, found in soil and the gastrointestinal tract of ruminants, humans and marine sponges. As a member of the genus '' Baci ...
'', ''
Staphylococcus aureus ''Staphylococcus aureus'' is a Gram-positive spherically shaped bacterium, a member of the Bacillota, and is a usual member of the microbiota of the body, frequently found in the upper respiratory tract and on the skin. It is often posit ...
'', ''
Escherichia coli ''Escherichia coli'' (),Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. also known as ''E. coli'' (), is a Gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus '' Esc ...
'', and ''
Pseudomonas aeruginosa ''Pseudomonas aeruginosa'' is a common encapsulated, gram-negative, aerobic– facultatively anaerobic, rod-shaped bacterium that can cause disease in plants and animals, including humans. A species of considerable medical importance, ''P. aer ...
'', with activity comparable to that of the antibiotic
ampicillin Ampicillin is an antibiotic used to prevent and treat a number of bacterial infections, such as respiratory tract infections, urinary tract infections, meningitis, salmonellosis, and endocarditis. It may also be used to prevent group B stre ...
. These results corroborate an earlier study that additionally reported antibacterial activity against ''Salmonella enterica'' serovar Typhimurium, ''
Streptococcus pyogenes ''Streptococcus pyogenes'' is a species of Gram-positive, aerotolerant bacteria in the genus '' Streptococcus''. These bacteria are extracellular, and made up of non-motile and non-sporing cocci (round cells) that tend to link in chains. They ...
'', and ''
Mycobacterium smegmatis ''Mycobacterium smegmatis'' is an acid-fast bacterial species in the phylum '' Actinomycetota'' and the genus '' Mycobacterium''. It is 3.0 to 5.0 µm long with a bacillus shape and can be stained by Ziehl–Neelsen method and the aurami ...
''. Extracts of the puffball have also been reported to have
antifungal An antifungal medication, also known as an antimycotic medication, is a pharmaceutical fungicide or fungistatic used to treat and prevent mycosis such as athlete's foot, ringworm, candidiasis (thrush), serious systemic infections such as cry ...
activity against ''
Candida albicans ''Candida albicans'' is an opportunistic pathogenic yeast that is a common member of the human gut flora. It can also survive outside the human body. It is detected in the gastrointestinal tract and mouth in 40–60% of healthy adults. It is usua ...
'', '' C. tropicalis'', ''
Aspergillus fumigatus ''Aspergillus fumigatus'' is a species of fungus in the genus '' Aspergillus'', and is one of the most common ''Aspergillus'' species to cause disease in individuals with an immunodeficiency. ''Aspergillus fumigatus'', a saprotroph widespread ...
'', '' Alternaria solani'', ''
Botrytis cinerea ''Botrytis cinerea'' is a necrotrophic fungus that affects many plant species, although its most notable hosts may be wine grapes. In viticulture, it is commonly known as "botrytis bunch rot"; in horticulture, it is usually called "grey mould" ...
'', and '' Verticillium dahliae''. A 2009 study found ''L. perlatum'' puffballs to contain
cinnamic acid Cinnamic acid is an organic compound with the formula C6H5-CH=CH- COOH. It is a white crystalline compound that is slightly soluble in water, and freely soluble in many organic solvents. Classified as an unsaturated carboxylic acid, it occurs n ...
at a concentration of about 14 milligrams per kilogram of mushroom. The fruit bodies contain the pigment
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amin ...
. The
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
lycoperdic acid (chemical name 3-(5(''S'')-carboxy-2-oxotetrahydrofuran-5(''S'')-yl)-2(''S'')-alanine) was isolated from the puffball, and reported in a 1978 publication. Based on the structural similarity of the new amino acid with (''S'')-
glutamic acid Glutamic acid (symbol Glu or E; the ionic form is known as glutamate) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can syn ...
, (''S'')-(+)-lycoperdic acid is expected to have
antagonistic An antagonist is a character in a story who is presented as the chief foe of the protagonist. Etymology The English word antagonist comes from the Greek ἀνταγωνιστής – ''antagonistēs'', "opponent, competitor, villain, enemy, ri ...
or
agonistic An agonist is a chemical that activates a receptor to produce a biological response. Receptors are cellular proteins whose activation causes the cell to modify what it is currently doing. In contrast, an antagonist blocks the action of the ag ...
activity for the
glutamate receptor Glutamate receptors are synaptic and non synaptic receptors located primarily on the membranes of neuronal and glial cells. Glutamate (the conjugate base of glutamic acid) is abundant in the human body, but particularly in the nervous syste ...
in the mammalian
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
. Methods to synthesize the compounds were reported in 1992, 1995, and 2002.


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{{featured article Lycoperdon Edible fungi Puffballs Fungi described in 1796 Fungi found in fairy rings Fungi of Asia Fungi of Australia Fungi of Central America Fungi of Europe Fungi of New Zealand Fungi of North America Fungi of Oceania Fungi of South America Taxa named by Christiaan Hendrik Persoon Fungi of Iceland